Selection for non-diapause inAmblyseius cucumeris andAmblyseius barkeri and exploration of the effectiveness of selected strains for thrips control

In Europe and North America the western flower thrips,Frankliniella occidentalis, is an important pest in various greenhouse crops, such as sweet pepper and cucumber. Two species of predatory mite are commercially applied for biological control of this pest:Amblyseius cucumeris andA. barkeri. Thrips control is generally successful from March onwards. During winter, however, thrips control by these predatory mites is less effective. An important reason for this is that the commercially applied strains of both mite species enter reproductive diapause under short-day photoperiods, whereas the western flower thrips does not enter diapause. In this paper we report on selection experiments for non-diapause in strains of both mite species, aimed at obtaining predators that do not enter diapause under light- and temperature conditions prevailing in winter. Additional experiments were done to estimate the potential of the selected lines as control agents ofF. occidentalis. Selection for non-diapause proved highly successful in both predatory mite species. In a New Zealand strain ofA. cucumeris diapause incidence decreased from 41% to 0% in about ten generations; in a Dutch strain ofA. barkeri diapause incidence decreased from 67% to 0% in about six generations. Furthermore, selection for non-diapause had no influence on predator performance, measured as predation rate and oviposition rate on a diet of first instar thirps larvae. Rates of predation and oviposition were the same for selected and unselected lines in both species; rates of predation and oviposition were higher forA. cucumeris than forA. barkeri. After 18 months under non-diapause conditions, no less than 92% of a sample of the selected non-diapause line ofA. cucumeris did not enter diapause when tested under diapause-inducing conditions. This indicates that ‘non-diapause’ is a stable trait in these predatory mites. Finally, a small-scale greenhouse experiment in a sweet pepper crop showed that the selected non-diapause line ofA. cucumeris established successfully under diapause-inducing short-day conditions.     

Evaluation of phytoseiid predators for control of western flower thrips on greenhouse cucumber

Ten predatory mite species, all phytoseiids, were evaluated for control of western flower thrips (WFT), Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), on greenhouse cucumber. This study was done to further improve biological control of thrips on this crop. Neoseiulus cucumeris (Oudemans) is at present used for biological control of thrips in greenhouses. Compared to this species, Typhlodromalus limonicus (Garman & McGregor), Typhlodromips swirskii (Athias-Henriot) and Euseius ovalis (Evans) reached much higher population levels resulting in a significantly better control of thrips. T. limonicus was clearly the best predator of WFT. Also Euseius scutalis (Athias-Henriot) increased to higher populations levels than N. cucumeris, but without controlling the thrips, probably because of an unequal distribution of this predator on the plant. Iphiseius degenerans (Berlese), Neoseiulus barkeri (Hughes), Euseius finlandicus (Oudemans) and Typhlodromus pyri (Scheuten) did not establish better than N. cucumeris. A non-diapausing exotic strain of N. cucumeris did not differ from the North European strain. The best performers in this study were all of sub-tropical origin. T. limonicus, T. swirskii and E. ovalis have good potentials for controlling not only thrips but also whiteflies. Factors affecting the efficacy of phytoseiids on greenhouse cucumbers are discussed.     

Cannibalism and Interspecific Predation in a Phytoseiid Predator Guild from Cassava Fields in Africa: Evidence from the Laboratory

Interspecific predation and cannibalism are common types of interaction in phytoseiid predator guilds, but the extent and nature of these interactions have not been determined yet in phytoseiid guilds composed of African native and neotropical exotic phytoseiid predators found in cassava habitat in southern Africa. We determined in laboratory experiments the level of cannibalism and interspecific predation among the three phytoseiid mite species Euseius fustis, Iphiseius degenerans, and Typhlodromalus aripo in the absence of food and in the presence of limited or abundant quantities of two food types – Mononychellus tanajoa and maize pollen – commonly found on cassava in Africa. When confined without food, only two T. aripo females laid each two eggs within 5 days, and this species survived longer than I. degenerans and E. fustis. In the presence of con- or hetero-specific larvae or protonymphs, the three species fed more on the former than on the latter, and more on hetero-specifics than on con-specifics. Oviposition rates of the three species did not exceed 0.7 egg/female/day on con- and hetero-specific immatures. Typhlodromalus aripo and E. fustis survived longer on con-specific and hetero-specific larvae and on hetero-specific protonymphs than in the absence of any food, while T. aripo survived longer than the two other species on the same diets. Provision of limited quantity of food decreased interspecific predation rate by I. degenerans and T. aripo, but not by E. fustis, and increased oviposition rate and longevity of all three species. Provision of abundant food, however, eliminated cannibalism by all three species and further reduced interspecific predation rates, but their oviposition and longevity remained relatively unchanged compared with limited food provision. Potential consequences of cannibalism and interspecific predation among phytoseiid mites on cassava for the biological control of M. tanajoa are discussed.     

ReleasingAmblyseius spp. [Acarina: Phytoseiidae] to controlThrips tabaci [Thysanoptera: Thripidae] on cabbage

Two species of predaceous mites,Amblyseius barkeri (Hughes) [=A. mckenziei Schuster & Pritchard] andA. cucumeris (Oudemans) [Acarina: Phytoseiidae] were evaluated as potential biological control agents forThrips tabaci Lindeman [Thysanoptera: Thripidae] on cabbage.A. barkeri colonized cabbage heads in preliminary trials. Field releases of different numbers of mites per plant during 1987 showed thatA. barkeri reduced the number of thrips in cabbage heads at harvest, and the reduction was proportional to the number of mites released, but thrips damage was unaffected. ReleasingA. cucumeris at different times during 1988, but releasing the same number of mites each time, we found that earlier releases resulted in fewer thrips and less damage at harvest; these relationships were not present, however, in plots treated with pyrethroid insecticides. We conclude that inoculative release ofAmblyseius spp. is a potentially useful thrips management strategy, but improvements in release timing and strategy will be required to provide commercially acceptable control.      

Predators induce egg retention in prey

To prevent predation on their eggs, prey often avoid patches occupied by predators. As a result, they need to delay oviposition until they reach predator-free patches. Because many species allocate energy to egg production in a continuous fashion, it is not clear what kind of mechanism prey use to delay oviposition. We used females of the phytoseiid mite Neoseiulus cucumeris to study these mechanisms. Females were placed in patches with pollen, a food source they use for egg production, and they were exposed to another phytoseiid mite, Iphiseius degenerans, which is an intraguild predator of N. cucumeris juveniles. We found that the oviposition of N. cucumeris females on patches with the predator was lower than on patches without the predator. Cues left by the intraguild predator were not sufficient to elicit such behaviour. Females of N. cucumeris reduced oviposition when exposed to the predator by retaining the egg inside their body, resulting in a lower developmental rate once these eggs were laid. Hence, females are capable of retaining eggs, but the development of these eggs continues inside the mother’s body. In this way, females gain some time to search for less risky oviposition sites.     

Influence of diet on life table parameters of <Emphasis Type="Italic">Iphiseius degenerans</Emphasis> (Acari: Phytoseiidae)

Studies on the reproduction, longevity and life table parameters of Iphiseius degenerans (Berlese) were carried out under laboratory conditions of 25 ± 1 °C, 75 ± 5% RH and 16L:8D h. As food sources for the predatory mite, Ricinus communis L. pollen, all stages of the spider mite Tetranychus urticae Koch, Frankliniella occidentalis (Pergande) larvae, and Ephestia kuehniella Zeller eggs were selected. All diets were accepted as food by the adult mites. Female longevity ranged from 29.5 to 42.4 days, the highest value was recorded on a diet of Ephestia eggs. The highest percentage of females escaping the experimental arena was observed on the diet consisting of thrips larvae. The highest oviposition rate (1.9 eggs/female.day) was recorded when the predator was fed on spider mites on an artificial substrate. For other diets, oviposition rates ranged from 1.0 to 1.3 eggs/female.day. The intrinsic rate of natural increase (r _m) of I. degenerans varied between 0.015 and 0.142 females/female.day. The diet consisting of castor bean pollen resulted in the highest population growth whereas the diet on spider mites brushed off onto a bean leaf arena resulted in the slowest population growth. This can be explained by the inability of the predator to cope with the webbing of T. urticae, and the high escape rate of the progeny when reared on spider mites. The percentage of females in the offspring ranged from 40 to 73%.This revised version was published online in May 2005 with a corrected cover date.     

The broad mite,Polyphagotarsonemus latus,as a potential prey for phytoseiid mites in California

In laboratory studies, individuals ofEuseius stipulatus Athias-Henriot,Typhlodromus rickeri Chant,T. porresi McMurtry andT. annectens DeLeon offered broad mite,Polyphagotarsonemus latus (Banks), as prey had developmental and ovipositional rates comparable to those offered a known favorable food (tetranychid mites or pollen).Euseius hibisci (Chant) had a longer developmental period and a markedly lower oviposition rate onP. latus than on the control food. Immature survival and ovipositional rates ofEuseius concordis Chant,Typhlodromus occidentalis Nesbitt,Amblyseius limonicus Garman and McGregor andIphiseius degenerans (Berlese) were very low onP. latus compared to control foods.     

Biological control of thrips [Thysanoptera: Thripidae] on greenhouse cucumber byAmblyseius cucumeris

The predatory miteAmblyseius cucumeris (Oudemans) [Acarina: Phytoseiidae] was evaluated as a biological control forThrips tabaci Lindeman andFrankliniella occidentalis (Pergande) [Thysanoptera: Thripidae] on greenhouse grown seedless cucumber.A. cucumeris spread throughout the greenhouses and provided control of both species of thrips.A. cucumeris adults persisted on plants for 7 weeks in the virtual absence of prey, and increased numerically in response to increases in prey population. On the basis of these resultsA. cucumeris is recommended as a useful biological control forT. tabaci andF. occidentalis on greenhouse cucumber. Publication No. 361, Agassiz Research Station, Agriculture Canada, Agassiz, B.C. Canada VOM 1AO.     

Effect of pollen,natural prey and factitious prey on the development of <Emphasis Type="Italic">Iphiseius degenerans</Emphasis>

The predatory mite Iphiseius degenerans (Berlese) is commercially available as a biological control agent of thrips and spider mites in greenhouse crops. Developmental duration and immature survival of I. degeneransreared on nine types of food (almond pollen, apple pollen, castor bean pollen, plum pollen, sweet pepper pollen, Tetranychus urticaeKoch, Frankliniella occidentalis(Pergande), Ephestia kuehniella Zeller eggs and Artemia franciscana Kellogg cysts) and on three substrates (Multicel, sweet pepper leaf, and bean leaf) were determined in the laboratory. All experiments were carried out at 25 °C. On Multicel, mean developmental times on pollen ranged from 6.0 to 7.1 days, with the lowest value recorded on almond pollen and the highest on sweet pepper pollen. When reared on castor bean pollen significantly longer developmental times were obtained on a sweet pepper leaf compared to a bean leaf or Multicel. Developmental duration when offered T. urticaeon Multicel ranged between 6.1 and 6.9 days, on a bean leaf development was completed in 8.0 days. The longest developmental times on Multicel were recorded on Ephestia eggs (7.0 days) and on decapsulated Artemia cysts (7.5 days). No development beyond the protonymphal stage occurred in the absence of food or when encapsulated Artemia cysts or thrips larvae were offered on Multicel. On a sweet pepper leaf and a bean leaf, respectively 6.7 and 10.0% of the eggs reached adulthood when thrips larvae were provided as food; developmental times recorded here, were 9.0 and 8.3 days. Overall, immature mortality occurred mainly in the protonymphal stage and ranged from 0.0 to 36.7%. In conclusion, I. degenerans is able to feed on a variety of natural and unnatural foods, but thrips larvae and sweet pepper pollen are unfavourable food for immature development. This could compromise the establishment of this biological control agent when used against thrips in sweet pepper crops.     

Nutritional benefits of intraguild predation and cannibalism among generalist and specialist phytoseiid mites

1. Intraguild predation and cannibalism are common among predaceous phytoseiid mites (Acari, Phytoseiidae) but the nutritional benefits gained by these processes are poorly understood. 2. The study reported here addressed the questions of whether cannibalism and intraguild predation provide different nutritional benefits and whether the ability to utilise cannibalism and intraguild predation is linked to the diet specialisation of phytoseiid mites. Specialists tested were Phytoseiulus macropilis, Galendromus occidentalis, and Neoseiulus longispinosus; generalists tested were Amblyseius andersoni, Neoseiulus cucumeris, and Neoseiulus fallacis. 3. All generalists and the specialist P. macropilis were able to complete juvenile development with both con‐ and hetero‐specific prey. Juvenile development of generalists was shorter with heterospecific prey than with conspecific prey, whereas development of the specialist P. macropilis did not differ between prey types. Only a few N. longispinosus and G. occidentalis, both specialists, reached adulthood by cannibalism but none reached adulthood by intraguild predation. 4. All generalists were able to sustain oviposition by intraguild predation. Neoseiulus cucumeris and A. andersoni laid more eggs with heterospecific prey than with conspecific prey, whereas N. fallacis had similar oviposition rates with both prey types. No specialist sustained oviposition by intraguild predation or cannibalism. 5. Overall, generalists gained equal or more nutritional benefits by intraguild predation than by cannibalism and were able to utilise phytoseiid intraguild prey as an alternative food source. Specialists gained equal or more nutritional benefits from cannibalism than from intraguild predation. For specialists, con‐ and hetero‐specific phytoseiids may be considered only a supplemental food.     

Functional response of Neoseiulus barkeri Hughes on two-spotted spider mite (Acari: Tetranychidae)

Generalist predatory mites are the common phytoseiid fauna in many agroecosystems, but little attention has been paid to their potential as biological control agents. In this study, we determined the functional responses of adult females of the generalist predator Neoseiulus barkeri Hughes on eggs, larvae, and adults of the two-spotted spider mite, Tetranychus urticae Koch, in the laboratory. Predation experiments were conducted on pepper leaf discs over a 24 h period at 25±1°C, 70–80% RH and 16L:8D photoperiod. Prey densities ranged 5 to 80 eggs, or 5 to 40 larvae, or 1 to 8 female adults of T. urticae per disc. The predation rate of N. barkeri adult females on T. urticae eggs was the same as on its larvae, but the predation rate on adult females was much lower. The role of generalist predatory mites in integrated and biological control of greenhouse pests was discussed.     

Effect of pollen supplement on intraguild predatory interactions between two omnivores: The importance of spatial dynamics

Arthropods often engage in complex trophic interactions such as intraguild predation (IGP), true omnivory (i.e., feeding on plants and prey), and apparent competition. Theoretical treatments of the effects of such interactions on herbivore populations have been concerned almost entirely with equilibrium conditions. Yet these interactions are common in non-equilibrium settings such as agroecosystems, where they are likely to have a strong influence on pest populations. We therefore tested short-term effects of IGP and food supplementation on interactions between two predators (the phytoseiid mite Neoseiulus cucumeris and the anthocorid bug Orius laevigatus) and their shared prey, Frankliniella occidentalis, on strawberry plants. All three consumers feed on strawberry pollen, both mites and bugs prey on thrips, and the bug also feeds on the mites (IGP). Strong IGP on mites (IG prey) by the bugs (IG predator) was recorded in structurally-simple arenas. In a more complex setting (whole-plants), however, the intensity of IGP differed among plant structures. Likewise, pollen supplementation reduced both IGP and predation on thrips in a structurally simple setting. In the whole-plant experiment, IGP was more intense on pollen-bearing than pollen-free flowers. The study illustrated how spatial dynamics, generated when consumers track food sources differently in the habitat and possibly when herbivorous and IG prey alter their distribution to escape predation, led to site-specific configuration of interacting populations. The intensity of resulting trophic interactions was weakened by food supplementation and by increased complexity of the habitat.     

Oviposition patterns in a predatory mite reduce the risk of egg predation caused by prey

Abstract 1. Predatory arthropods lay their eggs such that their offspring have sufficient prey at their disposal and run a low risk of being eaten by conspecific and heterospecific predators, but what happens if the prey attacks eggs of the predator? 2. The egg distribution and time allocation of adult female predatory mites Iphiseius degenerans as affected by predation of their eggs by prey, the western flower thrips Frankliniella occidentalis, were studied on sweet pepper plants. The predatory mites attack the first instar of thrips but all active stages of thrips are capable of killing the eggs of the predator; however the predatory mite is used for biological control of thrips. 3. The majority of predatory mite eggs was laid on the underside of leaves in hair tufts (domatia). During the experiment, females spent increasing amounts of time in flowers where they fed on pollen and thrips larvae. The risk of predation on predator eggs by thrips was lower on leaves than in flowers where the majority of thrips resides. Moreover, predation risk was higher outside leaf domatia than inside. 4. This suggests that predators avoid ovipositing in places with abundant prey to prevent their eggs from being eaten by thrips.     

Effects of high and low temperatures on development time and mortality of house dust mite eggs

The generalist predator Amblyseius swirskii is an efficient natural enemy of small insects and phytophagous mites, particularly thrips and spider mites. This phytoseiid species was considered for a long time as a subtropical species and Amblyseius rykei as a sub-Saharan African species. A recent revision of phytoseiid species of the subtribe Amblyseiina from sub-Saharan Africa Zannou et al. (Zootaxa 1550:1–47, 2007) determined that the two species are identical and synonymized them. To confirm or invalidate that morphological study, we crossed a Benin population of A. rykei and an Israel population of A. swirskii through two generations and back-crossed their hybrids to their parents. We also compared demographic parameters of both species on maize pollen, and their predation and oviposition rates on first larval instars of Frankliniella occidentalis. All females of homogamic and heterogamic crosses produced viable progeny, fertile F1 and viable F2. All the laid eggs hatched and sex ratio was female-biased for all crosses. Demographic parameters of the two species on maize pollen, and their predation rates and development times (egg to adult) on first instars of F. occidentalis were similar. Only oviposition of A. swirskii on larvae of F. occidentalis was significantly higher than that of A. rykei. These results indicate that A. rykei and A. swirkii are conspecific, and thus are a single species as concluded by Zannou et al.     

Diapause induction in the thrips predatorAmblyseius cucumeris [Acarina: phytoseiidae] under greenhouse conditions

The predatory miteAmblyseius cucumeris (Oudemans) is an important biological control agent for thrips in commercial greenhouses, but its effectiveness in fall and winter is limited by reproductive diapause induced under short day conditions. Influence of photoperiod and temperature on diapause induction was investigated to provide information for successful management of the predator. Under 8∶16 (L∶D) photoperiods and 22°C photophase temperatures, diapause incidence was inversely related to scotophase temperature, decreasing from 100% diapause at 15°C to no diapause at 21°C. In continuous darkness, diapause was induced by thermoperiods of 20∶10 and 22∶17 and 22∶17 but not 25∶15°C (T∶C) (8h thermophase). Critical daylength for inducing diapause under 22∶17°C (T∶C) was 12.45 h, which was consistent with the trend in diapause incidence in mites collected from an established greenhouse population September to November. MostA. cucumeris diapaused only when exposed to diapause inducing conditions throughout their juvenile development and none stopped laying eggs when transferred from nondiapause to diapause inducing conditions as adults. After 14 generations of genetic selection for a nondiapause strain, diapause incidence was 33.3%.      

Reproductive responses of Iphiseius degenerans and Neoseiulus teke (Acari: Phytoseiidae) to changes in the density of the cassava green mite, Mononychellus tanajoa (Acari: Tetranychidae)

We assessed the reproductive responses of adult female Iphiseius degenerans and Neoseiulus teke to increasing density of three stages of their prey, Mononychellus tanajoa, on cassava leaf discs under laboratory conditions. The oviposition rates increased with number of prey consumed per predator per day with a maximum of approximately two eggs per day for I. degenerans and four eggs per day for N. teke. The oviposition rate of N. teke was higher when consuming eggs than other prey stages. Neoseiulus teke was more efficient than I. degenerans in converting consumed prey into egg production. The data were adequately described by simple mathematical models.     

Factors affecting the distribution of a predatory mite on greenhouse sweet pepper

The predatory mite Neoseiulus cucumeris is used for biological control of phytophagous mites and thrips on greenhouse cucumber and sweet pepper. In a previous study, N. cucumeris provided effective control of broad mite but was only rarely found on the sampled leaves, raising questions about the factors affecting N. cucumeris distribution. To determine the distribution of N. cucumeris, leaves of pepper plants were sampled three times per day: just after sunrise, at noon and just before sunset for two years and throughout a 24 h period in one year. The presence of other mites and insects was recorded. Biotic (pollen) and abiotic (temperature, humidity) factors were monitored from the three plant levels. The effect of direct and indirect sunlight on the mites was assessed. N. cucumeris was found primarily in flowers; however, the mite’s distribution was affected by other predators (intraguild predation); in the presence of the predatory bug Orius laevigatus virtually no mites occurred in the flowers. Whereas temperature and humidity varied from the top to the lower level of the plants, apparently neither these factors nor the presence of pollen outside the flowers influenced mite distribution. N. cucumeris was found to be negatively phototropic; therefore N. cucumeris were pre-conditioned to light by rearing under light conditions for 4 months before being released. The light-reared mites were initially more numerous during the noon sampling period, however, rearing conditions caused only a temporary and non-significant change in distribution.     

Biological studies including population growth parameters of the predatory miteAmblyseius barkeri [Acarina.: Phytoseiidae] at 25°C in the laboratory

Amblyseius barkeri (Hughes) fed onThrips tabaci (Lind.) at 25°C showed an average duration of 2.2, 0.8 and 3.2 days for the egg, larval and nymphal stages, with mortalities at 1.0, 1.0 and 3.1%, respectively. Females represented 63% of the population and required multiple matings for optimal fertility. The oviposition period was 20.3 days and the average oviposition rate 2.3 eggs per day. The intrinsic rate of increase was 0.22 per day. The expected life span was 29.6 days for ♀♀ and 27.4 days for ♂♂.A. barkeri ♂♂ and ♀♀ both consumed 3.3 nymphs of thrips per day (mean value for the feeding stages). The larva does not take up food. In the absence of thripsA. barkeri was able to consume two-spotted spider mites,Tetranychus urticae (Koch), and their eggs, adult broad mites,Polyphagotarsonemus latus (Banks), and pollen of various plants. Cannibalism was observed when food was lacking. Certain morphological features, egglaying, mating and predatory behaviour are described.      

Intraguild predation among the predatory mites Amblyseius eharai,Amblyseius cucumeris and Amblyseius barkeri

Predator species with the same prey interact not only by competition for food and space but also by intraguild predation (IGP). The impact of IGP on introduced phytoseiid mites and native species in the context of biological control is a matter of considerable debate. Amblyseius eharai is the dominant native citrus species in central China, while Amblyseius cucumeris and Amblyseius barkeri are candidates for importation. All three species can feed on the spider mite Panonychus citri, which is the main pest in citrus. This study investigated, in the laboratory, possible IGP among these species in the absence and presence of P. citri, respectively. IGP in different densities of shared prey and intraguild (IG) prey was also studied. All three species consumed heterospecific larvae and eggs but not adults, and the IGP rate of larvae was significantly higher than that of eggs in the absence of shared prey. Additionally, the IGP rate of each group was reduced dramatically in the presence of both shared and IG prey when compared to the absence of shared prey. This occurs most likely because the three species prefer to feed on their natural prey P. citri, rather than on IG prey. Our results showed that A. eharai seems to be a more voracious IG predator than A. cucumeris. A. eharai was much more prone to IGP than A. barkeri.     

The benefits of clustering eggs: the role of egg predation and larval cannibalism in a predatory mite

Many arthropods produce clusters of eggs, but an unambiguous explanation for the evolution of egg clustering is still lacking. We test several hypotheses for the production of egg clusters by the predatory mite Iphiseius degenerans. This predator feeds on pollen, thrips larvae and nectar in flowers, but oviposits in clusters in tufts of leaf hairs (acarodomatia), where eggs run a lower risk of being killed by thrips, the prey of this predatory mite. The observed clustering is not caused by a shortage of oviposition sites; females preferably oviposit in a domatium containing eggs rather than in an empty domatium. To explain this preference, we first examined the effect of egg clusters on the risk of cannibalism. We found that eggs are invulnerable to cannibalism, whereas larvae emerging from single eggs or from clusters were equally vulnerable. Subsequently, we considered the killing of eggs resulting from counter-attacks by prey, i.e. the western flower thrips. We found no indication that a cluster of eggs protects eggs from predation by thrips. However, when eggs were clustered in a domatium rather than scattered over domatia, the proportion of eggs killed by thrips was lower. Hence, oviposition in clusters has no effect on its own and oviposition in domatia reduces predation risk by thrips, but oviposition in clusters in domatia leads to a synergistic effect on the survival of predator eggs. This synergism probably arises because eggs in clusters within tufts of leaf hairs are more difficult for thrips to reach. These experiments highlight a novel explanation of egg clustering, i.e. adaptation to counter-attacking prey. Moreover, they show that plant domatia protect predator eggs from predation.